38 research outputs found
Tracing the Domestic Pig Using the Omics Technologies
Pork represents one of the most important sources of protein in the human diet. Consumers today expect their food to be safe and of expected quality. Therefore, traceability and originality of the product must be guaranteed. This chapter provides an overview of the different approaches used for traceability and authentication of pork and pork products. Different DNA-based methods for meat speciation and authentication are described and their potential for use in the pork industry is highlighted
Estimation of population differentiation using pedigree and molecular data in Black Slavonian pig
Submitted 2020-07-17 | Accepted 2020-08-24 | Available 2020-12-01https://doi.org/10.15414/afz.2020.23.mi-fpap.241-249The aim of the study was to investigate the genetic differentiation of the Black Slavonian pig population. Two parallel analyses were performed using genealogical records and molecular data. Pedigree information of 6,099 pigs of the Black Slavonian breed was used to evaluate genetic variability and population structure. Additionally, 70 pigs were genotyped using 23 microsatellite markers. Genealogical data showed shrinkage in genetic diversity parameters with effective population size of 23.58 and inbreeding of 3.26%. Expected and observed heterozygosity were 0.685 and 0.625, respectively, and the average number of alleles per locus was 7.826. Bayesian clustering algorithm method and obtained dendrograms based on pedigree information and molecular data revealed the existence of four genetic clusters within the Black Slavonian pig. Wright’s FIS, FST and FIT from pedigree records were 0.017, 0.006, and 0.024, respectively, and did not prove significant population differentiation based on the geographical location of herds, despite the natural mating system. Obtained results indicate that despite the increased number of animals in the population, genetic diversity of Black Slavonian pig is low and conservation programme should focus on strategies aimed at avoiding further loss of genetic variability. Simultaneous use of genealogical and molecular data can be useful in conservation management of Black Slavonian pig breed.Keywords: autochthonous pig breed, microsatellite, genealogical data, genetic structuringReferencesBarros, E. A., Brasil, L. H. de A., Tejero, J. P., Delgado-Bermejo, J. V. & Ribeiro, M. N. (2017). Population structure and genetic variability of the Segureña sheep breed through pedigree analysis and inbreeding effects on growth traits. Small Ruminant Research, 149, 128-133.Belkhir, K. (2004). GENETIX 4.05, logiciel sous Windows TM pour la génétique des populations. http://www. genetix. univ-montp2. fr/genetix/genetix. htm.Boichard, D., Maignel, L. & Verrier, E. (1997). The value of using probabilities of gene origin to measure genetic variability in a population. Genetics Selection Evolution, 29, 5.Caballero, A. & Toro, M. A. (2000). Interrelations between effective population size and other pedigree tools for the management of conserved populations. Genetics Research, 75, 331-343.Casellas, J., Ibanez-escriche, N., Varona, L., Rosas, J. P. & Noguera, J. L. (2019). Inbreeding depression load for litter size in Entrepelado and Retinto Iberian pig varieties. Journal of Animal Science, 97(5), 1979–1986.Cortés, O., Martinez, A. M., Cañon, J., Sevane, N., Gama, L. T., Ginja, C., Landi, V., Zaragoza, P., Carolino, N., Vicente, A., Sponenberg, P. & Delgado, J. V. for the BioPig Consortium. (2016). Conservation priorities of Iberoamerican pig breeds and their ancestors based on microsatellite information. Heredity, 117(1), 14-24.Commission on Genetic Resources for Food and Agriculture Food and Agriculture Organization. (2011). Molecular genetic characterization of animal genetic resources. FAO.Croatian Agency for Agriculture and Food. (2020). Annual Report 2019: Pig breeding, Osijek, Croatia.Crovetti, A., Sirtori, F., Pugliese, C., Franci, O. & Bozzi, R. (2013). Pedigree analysis of Cinta Senese and Mora Romagnola breeds. Acta Agriculturae Slovenica, Suppl. 4, 41-44.D’Alessandro, E., Giosa, D., Sapienza, I., Giuffrè, L., Cigliano, R. A., Romeo, O. & Zumbo, A. (2019). Whole genome SNPs discovery in Nero Siciliano pig. Genetics and Molecular Biology, 42(3), 594-602.Diniz-Filho, J. A. F., Melo, D. B., de Oliveira, G., Collevatti, R. G., Soares, T. N., Nabout, J. C., Lima, J., Dobrovolski, R., Chaves, L. J., Naves, R. V., Loyola, R. D. & Telles M. P. de C. (2012). Planning for optimal conservation of geographical genetic variability within species. Conservation Genetics, 13(4), 1085-1093.Druml, T., Salajpal, K., Dikic, M., Urosevic, M., Grilz-Seger, G., & Baumung, R. (2012). Genetic diversity, population structure and subdivision of local Balkan pig breeds in Austria, Croatia, Serbia and Bosnia-Herzegovina and its practical value in conservation programs. Genetics Selection Evolution, 44(1), 5.Earl, D. A. & vonHoldt, B. M. (2012). STRUCTURE HARVESTER: a website and program for visualizing STRUCTURE output and implementing the Evanno method. Conservation Genetics Resources, 4(2), 359-361.Evanno, S., Regnaut, S. & Goudet, J. (2005). Detecting the number of clusters of individuals using the software STRUCTURE: A simulation study. Molecular Ecology, 14, 2611–2620.FAO (2000). Secondary guidelines for development of national farm animal genetic resources management plans: Management of small populations at risk. Rome: Food and Agriculture Organization.Francis, R. M. (2017). Pophelper: an R package and web app to analyse and visualize population structure. Molecular Ecology Resources, 17(1), 27-32.Goyache, F., Gutiérrez, J. P., Fernández, I., Gomez, E., Alvarez, I., Díez, J. & Royo, L. J. (2003). Using pedigree information to monitor genetic variability of endangered populations: the Xalda sheep breed of Asturias as an example. Journal of Animal Breeding and Genetics, 120, 95-105.Gutiérrez, J. P. & Goyache, F. (2005). A note on ENDOG: a computer program for analysing pedigree information. Journal of Animal Breeding and Genetics, 122, 172-176.Gvozdanović, K., Margeta, V., Margeta, P., Djurkin Kušec, I., Galović, D., Dovč, P. & Kušec, G. (2019). Genetic diversity of autochthonous pig breeds analyzed by microsatellite markers and mitochondrial DNA D-loop sequence polymorphism. Animal Biotechnology, 30(3), 242-251.Gvozdanović, K., Djurkin Kušec, I., Margeta, P., Salajpal, K., Džijan, S., Bošnjak, Z. & Kušec, G. (2020). Multiallelic marker system for traceability of Black Slavonian pig meat. Food Control, 109, 106917.International Society for Animal Genetics (ISAG)/Food and Agricultural Organization (FAO) (2011). Molecular genetic characterization of animal genetic resources. Rome: FAO Animal Production and Health Guidelines.Jombart, T. (2008). adegenet: a R package for the multivariate analysis of genetic markers. Bioinformatics, 24, 1403–1405.Jombart, T., Devillard, S. & Balloux, F. (2010). Discriminant analysis of principal components: A new method for the analysis of genetically structured populations. BMC Genetics, 11(1), 94.Kramarenko, S. S., Lugovoy, S. I., Kharzinova, V. R., Lykhach, V. Y., Kramarenko, A. S. & Lykhach, A. V. (2018). Genetic diversity of Ukrainian local pig breeds based on microsatellite markers. Regulatory Mechanisms in Biosystems, 9(2), 177-182.Lacy, R. C. (1987). Loss of genetic diversity from managed populations: interacting effects of drift, mutation, immigration, selection, and population subdivision. Conservation Biology, 1, 143-158.Lemus-Flores, C., Ulloa-Arvizu, R., Ramos-Kuri, M., Estrada, F. J. & Alonso, R. A. (2001). Genetic analysis of Mexican hairless pig populations. Journal of Animal Science, 79(12), 3021-3026.Lukić, B., Smetko, A., Mahnet, Ž., Klišanić, V., Špehar, M., Raguž, N. & Kušec, G. (2015). Population genetic structure of autochthonous Black Slavonian Pig. Poljoprivreda, 21(1), 28-32.Ma, L., Ya-Jie J. & Zhang, D. X. (2015). Statistical measures of genetic differentiation of populations: Rationales, history and current states. Current Zoology, 61(5): 886–897.Margeta, P., Margeta, V. & Budimir, K. (2013). How black is really Black Slavonian pig? Acta Agriculturae Slovenica, Suppl. 4, 25-28.Margeta, P., Margeta, V., Gvozdanović, K., Galović, D., Djurkin Kušec, I. & Kušec, G. (2016). Microsatellite multiplex method for potential use in Black Slavonian pig breeding. Acta Agriculturae Slovenica, 5, 66-70.Margeta, P., Gvozdanovic, K., Djurkin Kušec, I., Radišić, Ž., Kusec, G. & Margeta, V. (2018). Genetic analysis of Croatian autochthonous pig breeds based on microsatellite markers. Archivos de Zootecnia, 1, 13-16.Mariani, E., Summer, A., Ablondi, M. & Sabbioni, A. (2020). Genetic variability and management in Nero di Parma swine breed to preserve local diversity. Animals, 10(3), 538.Meuwissen, T. H. E. & Luo, Z. (1992). Computing inbreeding coefficients in large populations. Genetics Selection Evolution, 24, 305.Muñoz, M., Bozzi, R., García-Casco, J., Núñez, Y., Ribani, A., Franci, O., García, F., Škrlep, M., Schiavo, G., Bovo, S., Utzeri, V. J., Charneca, R., Martins, J. M., Quintanilla, R., Tibau, J., Margeta, V., Djurkin-Kušec, I., Mercat, M. J., Riquet, J., Estellé, J., Zimmer, C., Razmaite, V., Araujo, J. P., Radović, Č., Savić, R., Karolyi, D., Gallo, M., Čandek-Potokar, M., Fernández, A. I., Fontanesi, L. & Óvilo, C. (2019). Genomic diversity, linkage disequilibrium and selection signatures in European local pig breeds assessed with a high density SNP chip. Scientific Reports, 9(1), 13546.Nei, M. (1973). Analysis of gene diversity in subdivided populations. Proceedings of the National Academy of Sciences, 70(12), 3321-3323.Nei, M., Tajima, F. & Tateno, Y. (1983). Accuracy of estimated phylogenetic trees from molecular data. Journal of Molecular Evolution, 19(2), 153-170.Nei, M., (1987). Molecular Evolutionary Genetics. Columbia University Press, New York, 512 pp.Pritchard, J. K., Stephens, M. & Donnelly, P. (2000). Inference of population structure using multilocus genotype data. Genetics, 155, 945–959.Posta, J., Szabó, P. & Komlósi, I. (2016). Pedigree analysis of Mangalica pig breeds. Annals of Animal Science, 16(3), 701-709.R Development Core Team. (2018). A language and environment for statistical computing. R Foundation for Statistical Computing. Retrieved May 5, 2020 from http://www.R-project.org/.Sargolzaei, M., Iwaisaki, H. & Colleau, J. J. (2006). CFC: a tool for monitoring genetic diversity. Proc. 8th World Congr. Genet. Appl. Livest. Prod., CD-ROM Communication, (27-28), 13-18.Scali, M., Vignani, R., Bigliazzi, J., Paolucci, E., Bernini, A., Spiga, O., Niccolai, N. & Cresti, M. (2012). Genetic differentiation between CintaSenese and commercial pig breeds using microsatellite. Electronic Journal of Biotechnology, 15(2), 1-11.Silió, L., Barragán, C., Fernández, A.I., García‐Casco, J. & Rodríguez, M. C. (2016). Assessing effective population size, coancestry and inbreeding effects on litter size using the pedigree and SNP data in closed lines of the Iberian pig breed. 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Genetics Selection Evolution, 49(1), 71.Zhang, J., Jiao, T. & Zhao, S. (2016). Genetic diversity in the mitochondrial DNA D-loop region of global swine (Sus scrofa) populations. Biochemical and Biophysical Research Communications, 473(4), 814-820.
Black Slavonian (Crna slavonska) Pig
Black Slavonian (Crna slavonska) pig was created during the second part of the nineteenth century using planned crossing between four pig breeds. It is an autochthonous pig breed in the Republic of Croatia and one of the local pig breeds investigated in the project TREASURE. The present chapter aims to present history and current status of Black Slavonian pig breed, its exterior phenotypic characteristics, reproductive traits, geographical location, production system and main products from this breed of pigs. Also, a collection and review of available literature data, available until August 2017, on productive traits of Black Slavonian pig breed were carried out. Growth performance was estimated utilising average daily gain and average daily feed intake in the overall fattening stage as this was the information mostly provided in considered studies. Carcass traits were evaluated by means of age and weight at slaughter, hot carcass weight, carcass yield, muscularity and back fat thickness. Meat quality traits of the longissimus muscle evaluated were objective colour and intramuscular fat content. Although a considerable number of studies on Black Slavonian pig were included in the current review, data on growth performance and some parameters of carcass, meat and fat quality are scarce
COMPARISON OF COMMERCIAL DNA KITS AND TRADITIONAL DNA EXTRACTION PROCEDURE IN PCR DETECTION OF PORK IN DRY/FERMENTED SAUSAGES
In the present study four commercially available DNA extraction kits (Wizard® Genomic DNA Purification Kit, High Pure PCR Template Kit, DNeasy mericon Food and GeneJET PCR Purification Kit), as well as standard phenol/chloroform isolation technique have been evaluated regarding their concentration, purity and suitability for amplification of porcine DNA in dry/fermented sausages. The isolates were assessed for quantity and quality using spectrophotometer (IMPLEN GmbH, Germany). To verify template usability and quality of isolated DNA, the polymerase chain reaction (PCR) targeting at porcine cytochrome b by species specific primers was used. The comparison of extraction methods revealed satisfactory efficiency and purity of all extraction kits, while with standard phenol/chloroform isolation method high concentrations of DNA with low A260/280 were obtained. However, all the investigated techniques proved to be suitable for identification of porcine DNA in dry/fermented sausage. Thus, the standard phenol/chloroform DNA extraction method, as the cost-effective one, can be recommended as a good alternative to more expensive isolation kits when investigating the presence of pork DNA in dry/ fermented meat products
Verification of lean meat percentage estimation formulae for pig carcass classification in Croatia
Submitted 2020-07-23 | Accepted 2020-08-16 | Available 2020-12-01https://doi.org/10.15414/afz.2020.23.mi-fpap.265-268The study was performed on 136 pig carcasses representing the Croatian pig population with regards to the breed structure. The carcasses were dissected according to the EU reference method and lean meat percentage was estimated using the Hennessy Grading Probe (HGP7) device and the “Two points” (ZP) method. Comparison of lean meat percentage obtained by dissection and two prediction methods showed significant differences in estimating the lean meat percentage of pig carcasses (P <0.05). The distribution of carcasses according to SEUROP system showed a difference in the classification depending on the applied method, indicating a need for adjustment of current formulae for lean meat percentage estimation in Croatia.Keywords: pig, carcass, lean meat percentage, dissection, EU reference methodReferencesCauseur, D., Daumas, G., Dhorne, T., Engel, B., Fonti-Furnols, M., Hojsgaard, S. (2003). Statistical handbook for assessing pig classification methods: recommendations from the ‘EUPIGCLASS’ project group. EC working document.EEC Commission. (1994). Commission Regulation (EC) No 3127/94 of 20 December 1994 amending Regulation (EC) No 2967/85 laying Dowd detailed rules for the application of the Community scale for grading pig carcases. Official Journal of the European Communities, 43-44.EEC Commission. (2006). Commission Regulation (EC) No 1197/2006 of 7 August 2006 amending regulation (EEC) No 2967/85 laying down detailed rules for the application of the Community scale for grading pig carcasses. Official Journal of the European Union, 49, L 217, 8/8/2006, 6-7.EEC Commission. (2008). Commission Regulation (EC) No. 1249/2008 (2008, 12.16). of 10 December 2008 laying down detailed rules on the implementation of the community scales for the classification of beef, pig and sheep carcases and the reporting of prices thereof. Official Journal of the European Union, L337.EEC Commission. (2013). Regulation (EU) No. 1308/2013 (2013, 12.20). of the European Parliament and of the Council of 17 December 2013 establishing a common organisation of the markets in agricultural products and repealing Council Regulations (EEC) No 922/72, (EEC) No 234/79, (EC) No 1037/2001 and (EC) No 1234/2007. Official Journal of the European Union, L347.EEC Commission. (2017). Commission Implementng Regulaton (EU) 2017/1184 of 20 April 2017 laying down rules for the applicaton of Regulaton (EU) No 1308/2013 of the European Parliament and of the Council as regards the Union scales for the classifcaton of beef, pig and sheep carcasses and as regards the reportng of market prices of certain categories of carcasses and live animals. Official Journal of the European Union. htps://eur-lex.europa.eu/legal-content/EN/TXT/?uri= CELEX%3A32017R1184.Font-i-Furnols, M., Čandek-Potokar, M., Daumas, G., Gispert, M., Judas, M. & Seynaeve, M. (2016). Comparison of national ZP equations for lean meat percentage assessment in SEUROP pig classification. Meat science, 113, 1-8.Gangsei, L. E., Kongsro, J., Olsen, E. V., Røe, M., Alvseike, O. & Sæbø, S. (2016). Prediction precision for lean meat percentage in Norwegian pig carcasses using ‘Hennessy grading probe 7’: Evaluation of methods emphasized at exploiting additional information from computed tomography. Acta Agriculturae Scandinavica, Section A — Animal Science, 66(1), 17-24.Kušec, G., Kralik, G., Djurkin, I., Margeta, V., Maltar, Z. & Petričević, A. (2006). Comparison of different methods for lean percentage evaluation in pig carcasses. Acta Agraria Kaposváriensis, 10(2), 57-62.Kušec, G., Đurkin, I., Petričević, A., Kralik, G., Maltar, Z., Margeta, V. & Hanžek, D. (2007). Equations for lean share estimation in swine carcasses in Croatia. Poljoprivreda, 13 (1), 70-73.Kušec, G., Đurkin, I., Petričević, A., Kralik, G., Maltar, Z. & Margeta, V. (2009). Carcass leanness of pigs in Croatia estimated by EU referent method. Italian Journal of Animal Science, 8(3), 249-251.Kušec, G., Đurkin, I., Lukić, B., Radišić, Ž., Petričević, A. & Maltar, Z. (2011). The Equation for Prediction of Lean Meat Percentage by Hennessy Grading Probe in Croatia. Agriculturae Conspectus Scientificus, 76 (4), 329-331.NN 71/2018 Pravilnik o razvrstavanju i označivanju goveđih, svinjskih i ovčjih trupova te označivanju mesa koje potječe od goveda starih manje od 12 mjeseci.Sack, E. (1983). Using instruments to grade pork sides. Fleischwirtschaft, 63(3), 372–379.Vester-Christensen, M., Erbou, S.G.H., Hansen, M.F., Olsen, E.V., Christensen, L.B., Hviid, M., Ersboll, B.K. & Larsen, R. (2009). Virtual dissection of pig carcasses. Meat Science, 81, 699–704Walstra, P. & Merkus, G. S. M. (1996). Procedure for assessment of the lean meat percentage as a consequence of the new EU reference dissection method in pig carcass classification: based on discussion in the EU Management Committee on Pig Meat and based on discussions with dissection experts during a meeting on May 18-19, 1994 at Zeist, NL (No. 96.014). ID-DLO.
Carcass Composition and Meat Quality of Crna Slavonska Pigs from Two Different Housing Conditions
The aim of the study was to compare carcass composition and meat quality of pigs held in two housing systems suitable for production of Crna slavonska pigs. The experiment was conducted on 104 Crna Slavonska pigs (equal number of barrows and gilts) from two housing systems, outdoor (n=56) and indoor (deep bedding, n=48). The pigs in the outdoor group were reared until 18 months of age, and pigs in deep bedding group were raised until 15 months of age. After reaching the final age, the pigs were sacrificed and following carcass and meat quality traits were determined at the slaughter house and in laboratory: backfat and muscle thickness, length of the carcass from os pubis to atlas and from os pubis to 1st rib, ham length and circumference, loin eye area and fat area of longissimus lumborum muscle, pH values, water holding capacity and drip loss, colour reflectance scores (CIE L* a* b*) and instrumental tenderness. The carcasses of the pigs raised outdoors had lower backfat thickness, higher muscle thickness and lower fat area than pigs raised on deep bedding. Their carcasses were longer with longer hams. However, their ham circumference, as well as the loin eye area was lower than in pigs raised on deep bedding indicating higher production of muscle tissue in the latter. Out of investigated meat quality traits, the pigs raised outdoor had lower pH24 values measured in logissimus lumborum muscle and in semimembranosus muscle, higher drip loss and water holding capacity, higher cooking loss and yellowness (CIE b*)
Fatty acid composition of Baranjski kulen from two diverse production systems
Received: 2018-05-07 | Accepted: 2018-05-14 | Available online: 2018-11-26https://doi.org/10.15414/afz.2018.21.04.152-154The aim of the present study was to compare the fatty acid composition of dry fermented sausage made from Crna slavonska pig and modern hybrids reared in diverse production systems. The study was performed on Baranjski kulen, a traditional PGI (Protected Geographical Indication) labelled sausage. Baranjski kulen produced from pigs included in the study differed in their fatty acid profile, with kulen made from Crna slavonska pigs having a higher content of MUFA, lower content of PUFA and a more favourable PUFA/SFA ratio. The results of the present study demonstrate that meat originating from breeds raised in specific production system affects the fatty acid composition as one of the major determinants of nutritional product quality. Keywords: autochthonous breed, fatty acids, nutritional quality, pigsReferencesAgostoni, C., Moreno, L., Shamir, R. (2015) Palmitic Acid and Health: Introduction. Critical Reviews in Food Science and Nutrition, 12, 141-142. DOI: https://dx.doi.org/10.1080/10408398.2015.1017435Djurkin Kušec, I., Buha, I., Margeta, V., Gvozdanović, K., Radišić, Ž, Komlenić, M., Kušec, G. (2017) Carcass Composition and Meat Quality of Crna slavonska Pigs from Two Different Housing Conditions. Agriculturae Conspectus Scientificus, 82 (3), 221-225.Karolyi, D., Salajpal, K., Kiš, G., Đikić, M., Jurić, I. (2007) Influence of finishing diet on fatty acid profile of longissimus muscle of Black Slavonian pigs. Poljoprivreda, 13 (1), 176-179.Kasprzyk A., Tyra M., Babicz M. (2015) Fatty acid profile of pork from a local and a commercial breed. Archives Animal Breeding, 58, 379–385. DOI: https://dx.doi.org/10.5194/aab-58-379-2015Komlenić, M., Margeta, V., Djurkin Kušec, I., Gvozdanović, K., Margeta, P., Kušec, G. (2018) Carcass composition and meat quality of pigs from different pork chains in the production of Baranjski kulen (PGI). Archivos de Zootecnia, 209-212.Medić, H., Djurkin Kušec, I., Pleadin, J., Kozačinski, L., Njari, B., Hengl, B., Kušec, G. (2018) The impact of frozen storage duration on physical, chemical and microbiological properties of pork, Meat Science, 140, 119-127. DOI: https://dx.doi.org/10.1016/j.meatsci.2018.03.006Morse, N (2015). Are some health benefits of palmitoleic acid supplementation due to its effects on 5′ adenosine monophosphate-activated protein kinase (AMPK)? Lipid technology, 27 (12), 278-281. DOI: https://dx.doi.org/10.1002/lite.201500061Nevrkla, P., Kapelanski, W., Václavková, E., Hadaš, Z., Cebulska A., Horký, P. (2017) Meat quality and fatty acid profile of pork and backfat from an indigenous breed and a commercial hybrid of pigs. Annals of Animal Science, 17 (4), 1215-1227. DOI: https://dx.doi.org/10.1515/aoas-2017-0014Parunović, N., Radović, Č., Savić, R. (2017): Sensory properties and fatty acids profiles of fermented dry sausages made of pork meat from various breeds. IOP Conference Series: Earth and Environmental Sciences
Genotyping of the Leptin Receptor Gene in Crna Slavonska Pig – Preliminary Results Suggests New Variants of the Promoter
Researches on polymorphisms in the porcine LEPR gene and their association with economic traits were widely performed in the past. Manny polymorphisms in different part of the LEPR gene were described and majority of them was associated with economic traits such as growth and fatness. In present study, LEPR gene in 68 Crna Slavonska pigs was genotyped for HinfI polymorphism in the 3.8kb part of LEPR promoter, for HpaII and RsaI polymorphisms in the intron 4 and for ApeKI polymorphism in the exon 14. Allelic and genotype frequencies on polymorphic sites were calculated. Restriction of the 3.8 kb of the promoter region with HinfI revealed presence of two distinct restriction patterns, which haven’t been described so far. Their exact location and also their potential role in LEPR expression, as well as their impact on important economic traits should be explored in the future. Allelic and genotypic frequencies for other three polymorphic sites studied were more or less comparable with previous findings in the literature